By Calvin Victor Davis; Kenneth E Sorensen
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More extensive dis- cussion of these topics is to be found in the excellent book by Hodgkin (1964). A. Voltage Clamping. In Chapter 3, I characterized the action po- tential as an all-or-none process that continued to completion once the membrane potential has passed a threshold level. Since it was argued from experimental data that the apparent critical state variable is the membrane potential, and not membrane current, it is clear that a technique for placing the membrane potential under experimental control is essential for elucidating the processes underlying the generation of the action potential.
The experimenter sets the potential of the battery (actually a sophisticated electronic feedback system), and the battery supplies the current through the membrane necessary to maintain the desired potential. To tabulate the currents generally flowing through the membrane, it is necessary to extend the treatment of Chapter 2 slightly. membrane current, The jm' will be composed of two components - the dis- (0 VmI at) as before, and an ionic current m experimentally determined (not assumed to have placement current, c ji=ji(Vm,t) to be the ohmic behavior of Chapter 2).
This membrane current is identi- cally equal to the sum of the currents supplied from adjacent regions (r i +ro) -1 ( () 2Vml () x 2 ) and any current supplied from any external source, jc' such as the voltage clamp. 1) + j. 1 makes the principle of voltage clamping transparent. It is first experimentally arranged that ro is small (by, for example, placing the fibre in a large bath) and then inserting a wire inside the axon along its axis. The result of this procedure is to eliminate spatial variations in potential, effectively 'space clamping' the membrane by placing all of the membrane in direct electrical contact, with the resultant identical appearance of any membrane potential variation at all points on the axon.